The early phyllosoma stages of spiny echinatus Smith, 1869 (: Palinuridae) reared in the laboratory Abrunhosa, FA.a*, Santiago, AP.b* and Abrunhosa, JP.a aNúcleo de Estudos Costeiros, Campus de Bragança, Universidade Federal do Pará – UFPA, Alameda Leandro Ribeiro, Bairro Aldeia, CEP 68600-000, Bragança, PA, Brazil bLaboratório de Recursos Aquáticos, Universidade Federal do Ceará – UFC, Av. Mister Hull, s/n, Pici, CEP 60455-760, Fortaleza, CE, Brazil *e-mail: [email protected], [email protected] Received April 6, 2006 – Accepted September 1, 2006 – Distributed February 29, 2008 (With 6 figures)

Abstract The early stages of the Panulirus echinatus were hatched and reared in the laboratory. Ovigerous females were cap- tured in their habitat and carefully transported to the laboratory. Larvae were transferred in a recirculation water tank at a density of 10 larvae.L–1. The larvae were fed on Artemia and gonads of mussel Brachydonts sp. Microalgae Dunaliella viridis was added at a concentration of 150 x 104 cell.mL–1. Larvae and exuviae of each zoeal stage were preserved in an alcohol 70% + glycerin (1:1) solution. The phyllosomas moulted eight times; the intermoulting period of each instar averaged about 7 to 10 days. The main morphological changes of each appendage were described in detail, illustrated and compared with previous reports. Keywords: , phyllosoma, larval development, morphology.

Os primeiros estágios de filosoma da lagosta Panulirus echinatus (Decapoda: Palinuridae) cultivados em laboratório

Resumo Os estágios iniciais de lagosta tropical Panulirus echinatus eclodiram e foram cultivados em laboratório. Fêmeas ovígeras foram capturadas em seu habitat e cuidadosamente transportadas para o laboratório. As larvas foram cultiva- das em tanques de recirculação de água e alimentadas com náuplios de Artemia e gônadas mexilhão Brachydonts sp. Microalga Dunaliella viridis foi adicionada em cada tanque de cultivo na concentração de 150 x 104 cel.mL–1. Os filo- somas mudaram oito vezes e o período de intermuda de cada instar foi de cerca de 7 a 10 dias. As principais mudanças morfológicas de cada apêndice foram descritas, ilustradas e comparadas com trabalhos anteriores. Palavras-chave: lagosta, filosoma, desenvolvimento larval, morfologia.

1. Introduction The Brazilian spiny of the Panulirus genus lected in the sea (Lewis, 1951; Baisre, 1964; Phillips and have high economic value. The species P. argus (Latreille, Sastry, 1980) and for those reared in the laboratory pro- 1804) and P. laevicauda (Latreille, 1817) are captured gressed through the metamorphic puerulus stage (Inoue, extensively, representing one of the most important fish- 1981; Kittaka and Kimura, 1989; Yamakawa et al., 1989; ery resources in the world (Phillips et al., 1980). For a Matsuda and Yamakawa, 2000). The long larval period long time another species, P. echinatus, was occasionally (about one year) is considered an important obstacle for captured, however, an increase in their production has phyllosoma culture research. The lack of detailed morpho- been observed (Santiago, 2001). Despite the economic logical description of some or all phyllosoma stages has significance that this latter species has represented in the also hindered studies of larval distribution and ecology. fisheries, few studies concerning their biological aspects Successful studies on culture from egg to the pueruli have been reported. stage of Panulirus were achieved for P. ­japonicus (Von Information on the biology of phyllosomas is consid- Siebold, 1824) (Kittaka and Kimura, 1989; Yamakawa ered important and necessary to elucidate aspects related et al., 1989) and P. longipes (A. Milne-Edwards, 1868) to larval biology and culture. The larval development (Matsuda and Yamakawa, 2000). Some attempts at rear- of spiny lobster has been reported for specimens col- ing phyllosoma larvae were reported for P. homarus

Braz. J. Biol., 68(1): 179-186, 2008 179 Abrunhosa, FA., Santiago, AP. and Abrunhosa, JP.

(Linnaeus, 1758) (Radhakrishnan and Vijayakumaran, gonads of Brachyodonts sp. Microalgae Dunaliella 1995) and for P. ornatus (Fabricius, 1776) (Duggan viridis Teodoresco 1906 were added daily in the culture and McKinnon, 2003) in which, 10 and 9 early instars system. were reared and described, respectively. More recently, Samples of larvae were removed after each instar Abrunhosa et al. (2004) described and compared the and fixed in 10% formalin solution for 24 hours and morphological characteristics of three early instars of P. preserved in a glycerol+alcohol 70% (1:1) solution. The laevicauda and P. echinatus, contributing to the identifi- larvae were dissected under an optical binocular micro- cation of phyllosomas that occur on the Brazilian coast. scope. The illustrations of each larval instar, including These three early larval instars of P. echinatus are again appendages, were made using a camera lucida. reviewed in the present study. For the species P. homarus, information on the av- The present study describes in detail and illustrates erage length, number of setae in the apical segment of the 5 early stages and instars of each stage of the spe- the 2nd maxilla and number of the exopod setae of each cies P. echinatus in the laboratory. Morphological com- appendage were based on the illustrations reported by parisons with larvae of its congeneric species, previously Radhakrishnan and Vijayakumaran (1995). described, are briefly discussed. The terminologies STAGES and INSTAR were used in order to make comparisons with other previously de- 2. Material and Methods scribed species. Both terms were extensively used by Ovigerous females of P. echinatus were obtained at the authors in descriptions of larval development of the Iparana Beach (Caucaia/CE) in October 2001 by autono- spiny lobster. mous diving in the coastal shallow waters. They were transported to the laboratory in plastic baskets (10 L) 3. Results containing local seawater. In the laboratory the females were acclimated and 3.1. On the larval culture then placed in a circular tank (cap. 1000 L) with biologi- The larval hatching was closely related to the moon cal filter and constant aeration. After hatching, the phyllosomas were carefully phases, in which the majority of the larval hatching was transferred to three transparent water recirculation tanks carried out on the first quarter and full moon, indicating at a density of 60 larvae.L–1. These tanks are similar to that lunar cycle had direct influence in the life cycle in those used for culture of many species of spiny lobsters the laboratory. of the family Palinuridae (Kittaka, 1994). The tanks, The phyllosomas readily feed on Artemia nauplii which were confectioned with acrylic (70 cm Ø and in stages I to III. Then, the phyllosomas seemed to 30 cm deeper), allow free circulation of flowing water have difficulty in catching Artemia so other foods were and total visualization of the floating elements is pos- tested. Some foods were initially well accepted by the sible. Salinity was maintained at 35‰. The larvae were ­phyllosomas, but later were rejected. This behaviour fed Artemia nauplii for the 3 first instars and then, with may be related to nutritional aspects, because many

Table 1. Intermoulting period, accumulative days and maim characteristics of each stage for phyllosomas of Panulirus ­echinatus reared in the laboratory. Stage Intermoult Accumulative Main characteristics (Mean days) days I 9 9 Ocular peduncle unsegmented. Exopod of the 1st and 2nd pereiopods with 5 pairs of setae. II 9 18 Eyes segmented; exopod of the 1st and 2nd pereiopods with 6 pairs of setae. IIIa 7 25 Fourth pereiopod present; exopod of the 1st and 2nd pereiopods with 7 pairs of setae. IIIb 5 30 1st and 2nd pereiopods with 8 pairs of setae. IVa 5 35 Fifth pereiopod present; exopod of the 1st and 2nd ­pereiopods with 9 pairs of setae. IVb 5 40 exopod of the 1st and 2nd pereiopods with 10 pairs of setae. IVc 7 47 exopod of the 1st and 2nd pereiopods with 11 pairs of setae. IVd 8 55 exopod of the 1st and 2nd pereiopods with 12 pairs of setae. V >8* >63 Antennule 3-segmented; exopod of the 1st and 1st and 2nd pereiopods with 13 pairs of setae. *No phyllosoma succeeded in moulting to the next instar.

180 Braz. J. Biol., 68(1): 179-186, 2008 The early phyllosoma stages of spiny lobster

­phyllosomas failed to moult to the subsequent stage. This First Maxilla (Figure 2c): Coxal endite with 2 long fact was more evident for larvae in the VIII instar. serrulate setae and 1 short simple seta; basal endite with The larvae moulted eight times, once at stages I to II, 2 strong serrulate setae distally and 1-2 sub-distally; en- two times at stage III and four times at stage IV (Table 1). dopod short, unsegmented with 2 setae. There was not a specific day or night period in which Second Maxilla (Figure 2d): Basal segment en- the larvae moulted to the next instar. The moult of the larged with 2-3 short setae, distal segment short with phyllosomas from stage I to II and II to III ranged from 7 4 long plumose setae. to 11 and 7 days, averaging in 9 and 7 days, respectively. First Maxilliped: Small, uniramous and unsegment- For successive instars, it was not possible to determine ed protuberance. accurately the intermoult period for later instars, because Second Maxilliped (Figure 2e): Endopod 5-seg- the phyllosomas were not reared individually. The inter- mented, distal segment ending in 1 strong distal and moult average for each instar is shown in Table 1. 2 sub-distal spines, sub-distal segment rounded distally The largest mortality of the phyllosomas was on the with long serrulate setae; exopod absent. th 4 day of rearing, when about 50% of the larvae died in Third Maxilliped: Endopod well developed, 5-seg- both systems. After this period, low mortality rates were mented, segment distal with long setae rounding the observed. distal margin and 2 long setae on the sub-distal margin; 3.2. On gross morphology exopod with 3 pairs of long and plumose setae. The morphological features were fully described First and second Pereiopods: Endopod 5-segment- only for instar I. For the subsequent instars, only main ed, completely developed, distal segment ending in a morphological changes were reported. The morphologi- strong spine, dorsal and ventral sub-exopodal spines ab- cal changes and the main characteristics of each stage are sent, coxal spine present but not elongated (disposition fully described in the Table 2. of setae as illustrated); exopod with 5 pairs of long and plumose setae. STAGE I with one instar (Figure 1a) Third Pereiopod: Endopod very long, 5-segmented, ending in a strong curved spine, coxal spine present; Cephalic shield: Mean length 1.04 mm (range: exopod undeveloped showing a small unarmed protuber- 1.00-1.07), semicircular, pear like in shape. ance. Eyes: Ocular peduncle unsegmented. Abdomen: Short, unsegmented with 3 setae and Antennule (Figure 2a): Uniramous, unsegmented, 1 spine at each ramous on the posterior portion. longer than antenna, ending in a short spine, 3 long aes- thetascs and 2 simple setae. STAGE II with one instar (Figure 1b) Antenna (Figure 2b): Uniramous, 2-incompletely segmented, ending in a short curved spine and 2-3 sim- Cephalic shield: Mean length 1.25 mm (range: ple setae, 2 simple setae in the distal region near to the 1.18-1.30 mm). apex. Eyes: Ocular peduncle segmented.

Table 2. Cephalic length, number of setae of the 2nd maxilla and exopod of maxillipeds and pereiopods of Panulirus ­echinatus reared in the laboratory. Stage Instar Average of 2nd maxilla Pairs of setae in the exopods Cephalic (Distal seg. ) 3rd 1st 2nd 3rd 4th 5th shield Length maxill. Pereiop. Pereiop. Pereiop. Pereiop. Pereiop. (mm) I I 1.04 4 3 5 5 0 absent absent II II 1.25 4 3 6 6 0 absent absent III III 1.55 4 4 7 7 3 bud absent IV 1.72 4 4 8 8 4 bud absent IV V 2.09 4 5 9 9 5 bud bud birram. VI 2.17 4 5 10 10 6 bud bud birram. VII 2.25 4 6 11 11 7 bud bud birram. VIII 2.50 4 7 12 12 8 0 bud V* IX 2.58 4 8 13 13 9 0 bud *The culture finished at the first instar of the stage V (nine instars were reared.)

Braz. J. Biol., 68(1): 179-186, 2008 181 Abrunhosa, FA., Santiago, AP. and Abrunhosa, JP.

a an1

Template de Figuras - BJB an2 mp3 * Fontes Times (Roman), tamanho 8. * "Cenário" - linhas com 0.5 de Stroke. mp2 p1 mx2 b * Linhas pertencentes a "Dados gráficos" commp1 0.6 de Stroke. * Sempre que houver rosa do ventos na imagem original, substituir pelacs padrão da paleta symbol. an1 * Preenchimento de barras com 10% de preto quando houver texto e 50% quando não. mp3 * Dados na tabela ou figura devem estar todos em Inglês. p3 an2 * Legendas devem estar dentro de caixas de texto com 2 mm de distância nas extremidades. * Texto da figura ou gráfico deve estar em "Sentence case". mp2 mx2 * Escala de mapas deverá ter 1 ponto de Stroke. p3 mp1 * Letras que representam figuras ex: a , devem estar no canto superior direito com 2 mm de distância das extremidades da figura. cs pe1

pe3 pe2

0.5 mm

0.5 mm

Figure 1. Panulirus echinatus (in ventral view): a) Stage I; b) Stage II. Abbreviations: an1 = antennule; an2 = antenna; cs = coxal spine; mp1 = 1st maxilliped; mp2 = 2nd maxilliped; mp3 = 3rd maxilliped; mx2 = maxilla; pe1 = 1st pereiopod; pe2 = 2nd pereiopod; pe3 = 3rd pereiopod.

a b

d

c e

Figure 2. Panulirus echinatus: a) antennule, b) antenna, c) maxillule, d) maxilla, and e) 2nd maxilliped. Scale bar = 0.5 mm.

182 Braz. J. Biol., 68(1): 179-186, 2008 The early phyllosoma stages of spiny lobster

First and second Pereiopods: Exopod with 6 pairs Third Maxilliped: Exopod with 5 pairs of long and of long and plumose setae. plumose setae. Third Pereiopod: Exopod a little more developed First and second Pereiopods: Exopod with 9 pairs compared to the first instar, lacking setae. of long and plumose setae. Third Pereiopod: Exopod with 5 pairs of long and STAGE III with two instars plumose setae. Fourth Pereiopod: More developed in relation to INSTAR IIIa (Figure 3a) previous instar (as illustrated), lacking setae. Cephalic shield: Mean length 1.55 mm (range: Fifth Pereiopod: Present but unsegmented, bud and 1.38-1.74 mm). lacking setae. Third Maxilliped: Exopod with 4 pairs of long and plumose setae. INSTAR IVb (Figure 4b) First and second Pereiopods: Exopod with 7 pairs Cephalic shield: Mean length 2.17 mm (range: 2.05- of long and plumose setae. 2.39 mm). Third Pereiopod: Exopod well developed compared First and second Pereiopods: Exopod with 10 pairs to previous instars with 3 pairs of long and plumose se- of long and plumose setae. tae. Third Pereiopod: Exopod with 6 pairs of long and Fourth Pereiopod: Unsegmented and biramous plumose setae. bud. INSTAR IVc (Figure 5a) INSTAR IIIb (Figure 3b) Cephalic shield: Mean length 2.25 mm (range: Cephalic shield: Mean length 1.72 mm (range: 2.15-2.35 mm). 1.70-1.75 mm). Third Maxilliped: Exopod with 6 pairs of long and First and second Pereiopods: Exopod with 8 pairs plumose setae. of long and plumose setae. First and second Pereiopods: Exopod with 11 pairs Third Pereiopod: Exopod with 4 pairs of long and of long and plumose setae. plumose setae. Third Pereiopod: Exopod with 7 pairs of long and plumose setae. STAGE IV with 4 instars INSTAR IVd (Figure 5b) INSTAR IVa (Figure 4a) Cephalic shield: Mean length 2.50 mm (range: Cephalic shield: Mean length 2.09 mm, narrower 2.48-2.52 mm). than previous instars, about 1/3 longer than large (range: Antennule: Segmented, proximal segment longer 1.90-2.28 mm). than the distal one.

Template de Figuras - BJB a

* Fontes Times (Roman), tamanho 8. an1 Template de Figuras - BJB b * "Cenário" - linhas com 0.5 de Stroke. an1 * Linhas pertencentes a "Dados gráficos" com 0.6 de Stroke. * Fontes Times (Roman), tamanho 8. an2 * Sempre que houver rosa do ventos na imagem original, substituir pela* "Cenário" padrão da - linhaspaleta comsymbol. 0.5 de Stroke. an2 mp3 mp3 * Preenchimento de barras com 10% de preto quando houver texto e *50% Linhas quando pertencentes não. a "Dados gráficos" com 0.6 de Stroke. * Dados na tabela ou figura devem estar todos em Inglês. * Sempre que houver rosa do ventos na imagem original, substituir pela padrão da paleta symbol. mp2 * Legendas devem estar dentro de caixas de texto com 2 mm de distância* Preenchimento nas extremidades. de barras com 10% demp2 preto quando houver texto e 50%pe1 quando não. mx2 mx2 mp1 * Texto da figura ou gráfico deve estar em "Sentence case". * Dados na tabela ou figura devem estar todos em Inglês. mp1 pe1 * Escala de mapas deverá ter 1 ponto de Stroke. * Legendas devem estar dentro de caixas de texto com 2 mm dece distânciape2 nas extremidades. * Letras que representam figuras ex: a , devem estar no canto superior* Texto direito da com figura 2 mmou gráfico de deve estar em "Sentence case". distância das extremidades da figura. * Escala de mapas deverá ter 1 ponto de Stroke. pe4 pe2 pe3 * Letras que representam figuras ex: a , devem estarpe4 no canto superiorpe3 direito com 2 mm de distância das extremidades da figura.

1.0 mm

0.5 mm

Figure 3. Panulirus echinatus (in ventral view): a) Stage IIIa, b) Stage IIIb. Abbreviations: an1 = antennule; an2 = antenna; cs = coxal spine; mp1 = 1st maxilliped; mp2 = 2nd maxilliped; mp3 = 3rd maxilliped; mx2 = maxilla; pe1 = 1st pereiopod; pe2 = 2nd pereiopod; pe3 = 3rd pereiopod; pe4 = 4th pereiopod.

Braz. J. Biol., 68(1): 179-186, 2008 183 Abrunhosa, FA., Santiago, AP. and Abrunhosa, JP.

Template de Figuras - BJB an1 an1 a b * Fontes Times (Roman), tamanho 8. Template de Figuras - BJB * "Cenário" - linhas com 0.5 de Stroke. mp3 * Fontes Times (Roman), tamanho 8. an2 an2 * Linhas pertencentes a "Dados gráficos" com 0.6 de Stroke. * "Cenário" - linhas com 0.5 de Stroke. * Sempre que houver rosa do ventos na imagem original, substituir pela padrão da paleta symbol. * Linhas pertencentes a "Dados gráficos" com 0.6 de Stroke. * Preenchimento de barras com 10% de preto quando houver texto e 50% quando não. mp2 mp2 * Sempre que houver rosa do ventos na imagem original, substituir mp3pela padrão da paleta symbol. * Dados na tabela ou figura devem estar todos em Inglês. mx2 mx2 pe1 * Preenchimento de barras com 10%mp1 de preto quando houver texto e 50% quando não. mp1 * Legendas devem estar dentro de caixas de texto com 2 mm de distância nas extremidades. * Dados na tabela ou figura devem estar todos em Inglês. pe1 * Texto da figura ou gráfico deve estar em "Sentence case". * Legendas devem estar dentro de caixas de texto com 2 mm de distância nas extremidades. * Escala de mapas deverá ter 1 ponto de Stroke. pe2 * Texto da figura ou gráfico deve estar em "Sentence case". * Letras que representam figuras ex: a , devem estar no canto superior direito com 2 mm de pe4 pe4 pe5 pe2 pe5 pe3 distância das extremidades da figura. * Escala de mapas deverá ter 1 ponto de Stroke. pe3 * Letras que representam figuras ex: a , devem estar no canto superior direito com 2 mm de distância das extremidades da figura.

1.0 mm 1.0 mm

Figure 4. Panulirus echinatus (in ventral view): a) Stage IVa, b) StageIVb. Abbreviations: an1 = antennule; an2 = antenna; mp1 = 1st maxilliped; mp2 = 2nd maxilliped; mp3 = 3rd maxilliped; mx2 = maxilla; pe1 = 1st pereiopod; pe2 = 2nd pereiopod; pe3 = 3rd pereiopod; pe4 = 4th pereiopod; pe5 = 5th pereiopod.

b Template de Figuras - BJB an1 a

* Fontes Times (Roman), tamanho 8. an1 * "Cenário" - linhas com 0.5 de Stroke. an2 an2 * Linhas pertencentes a "Dados gráficos" com 0.6 de Stroke. mp3 * Sempre que houver rosa do ventos na imagem original, substituir pela padrão da paleta symbol. mp3 * Preenchimento de barras com 10% demp2 preto quando houver texto e 50% quando não. * Dados na tabela ou figura devem estarmx2 todos em Inglês. mp2 mp1 pe1 pe1 mx2 * Legendas devem estar dentro de caixas de texto com 2 mm de distância nas extremidades. mp1 * Texto da figura ou gráfico deve estar em "Sentence case". * Escala de mapas deverá ter 1 ponto de Stroke. * Letras que representam figuras ex: a , devem estar no canto superior direito com 2 mm de pe2 pe5 pe4 distância das extremidades da figura. pe3 pe2 pe5 pe4 pe3

1.0 mm 1.0 mm

Figure 5. Panulirus echinatus (in ventral view): a) Stage IVc, b) Stage IVd. Abbreviations: an1 = antennule; an2 = antenna; mp1 = 1st maxilliped; mp2 = 2nd maxilliped; mp3 = 3rd maxilliped; mx2 = maxilla; pe1 = 1st pereiopod; pe2 = 2nd pereiopod; pe3 = 3rd pereiopod; pe4 = 4th pereiopod; pe5 = 5th pereiopod.

Third Maxilliped: Exopod with 7 pairs of long and STAGE V (Figure 6) plumose setae. First and second Pereiopods: Exopod with 12 pairs Cephalic shield: Mean length 2.58 mm (range: of long and plumose setae. 2.50-2.58 mm). Third Pereiopod: Exopod with 8 pairs of long and Antennule: 3-segmented, about 2 times the anten- plumose setae. nal size. Fourth Pereiopod: More developed in relation to previous instar (as illustrated), surpassing abdomen. Third Maxilliped: Exopod with 8 pairs of long and Exopod more developed, lacking setae. plumose setae.

184 Braz. J. Biol., 68(1): 179-186, 2008 The early phyllosoma stages of spiny lobster

other reported Panulirus species such as ­P. ­laevicauda Template de Figuras - BJB an1 (by Abrunhosa, 2004), P. ornatus (by Duggan and McKinnon, 2003) P. japonicus (Inoue, 1981), * Fontes Times (Roman), tamanho 8. ­P. ­longipes (by Matsuka and Yamakawa), ­­­­P. ­­h­o­marus (by * "Cenário" - linhas com 0.5 de Stroke. an2 Radhakrishnan and Vijayakumaran, 1995) and ­P. ­inflatus * Linhas pertencentes a "Dados gráficos" com 0.6 de Stroke. (by Inoue, 1981). The initial number of setae 3, 5, 5 in * Sempre que houver rosa do ventos na imagem original, substituir pela padrão da paleta symbol. mp3 the exopods of the third maxilliped and pereopods and * Preenchimento de barras com 10% de preto quando houver texto e 50% quando não. their gradual increase are also similar to the reported spe- * Dados na tabela ou figura devem estar todos em Inglês. mp2 cies. mx2 * Legendas devem estar dentro de caixas de texto com 2 mm de distância nas extremidades. mp1 pe3 On the other hand, the distinct difference among oth- * Texto da figura ou gráfico deve estar em "Sentence case". er species is the presence of coxal spines on P. e­chinatus, * Escala de mapas deverá ter 1 ponto de Stroke. which is included in group B, according to Gurney, 1936 pe2 * Letras que representam figuras ex: a , devem estar no canto superior direito com 2 mm de and Baisre and Ruiz de Quevedo, 1982. According to distância das extremidades da figura. pe5 pe3 Abrunhosa et al. (2004), phyllosomas of the Brazilian species, P. echinatus and P. argus, bear only the coxal pe4 spines while P. laevicauda exhibited both the coxal and sub-exopodal spines placing these species in different groups. 1.0 mm The Intermoult period appears to be distinct for pal- inurid species and may vary in accordance to the rearing Figure 6. Panulirus echinatus (in ventral view): Stage V. Abbreviations: an1 = antennule; an2 = antenna; mp1 = conditions in which the larvae are submitted, such as, ad- 1st maxilliped; mp2 = 2nd maxilliped; mp3 = 3rd maxilliped; aptation, type of food administered, density, water qual- mx2 = maxilla; pe1 = 1st pereiopod; pe2 = 2nd pereiopod; ity, and others (Kittaka and Ikegani, 1988; Kittaka et al., pe3 = 3rd pereiopod; pe4 = 4th pereiopod; pe5 = 5th pereio- 1988; Kittaka and Ikegami, 1988; Kittaka and Kimura, pod. 1989; Radhakrishnan and Vijayakumaran, 1995; Kittaka et al., 1998; Matsuda and Yamakawa, 2000; Sekine et al., 2000). Kittaka and Kimura (1989) rearing phyllosomas of First and second Pereiopods: Exopod with 13 pairs P. japonicus, obtained an intermoult period of 6 to 7 days of long and plumose setae. for instar I. However, a gradual increase in that period Third Pereiopod: Exopod with 9 pairs of long and was observed for the subsequent instars. Matsuda and plumose setae. Yamakawa (2000) observed an intermoult period for the Fourth Pereiopod: Surpassing about 2 times the first two instars of P. longipes similar to P. echinatus of abdomen. 7 to 10 days. But, for P. homarus, cultured individually, Fifth Pereiopod: Still bud and lacking setae. this period increased to 8 to 10 days. When P. homarus were submitted to mass culture, this period increased to 4. Discussion 12 to 14 days (Radhakrishnan and Vijayakumaran, 1995). In the laboratory, phyllosoma larvae moulted eight These facts may be related to the elevated density. times comprising five larval stages considering the Feeding is considered a critical factor in the suc- developmental criteria of description adopted by the cess of the phyllosoma culture. Delay or decrease of the authors in previous studies, Inoue (1978, 1981) for amount of food requested by the larva may prolong the P. ­japonicus, Radhakrishnan and Vijayakumaran (1995) intermoulting period or cause death during the ecdyse for P. homarus , Matsuda and Yamakawa (2000) for (Abrunhosa and Kittaka, 1997). High mortality was ob- P. longipes and Coutures, (2000) for palinurid species. served in the instar IV of P. echinatus. This fact may be The term instar, which corresponds to each moulting related to the nutritional requirement needed for moult- of the phyllosoma, was used for palinurid and scyllarid ing as many phyllosomas failed to moult to the subse- species by Kittaka (1994), Booth and Phillips (1994) and quent stage. An interesting study on larval feeding of Kittaka and Abrunhosa (1997). In general, the number P. elephas revealed their ability to capture large-sized of setae increased in the exopods of the phyllosoma ap- fish larvae (Kittaka and Abrunhosa, 1997). Presumably, pendages. In the case of P. echinatus, an additional pair of fish larvae may compose the natural diet for phyllosomas setae arises on each successive larval moulting (Table 2). in the sea. Such morphological change appears to be a characteris- Many kinds of food were tested in phyllosoma tanks tic of the palinurid species, in which each instar could be from instar III. Larvae of P. japonicus were cultured be- considered a larval stage. In this case, comparisons with ing fed with adult Artemia and pieces of mussel gonads other species will become easier to observe. (Sekine et al., 2000). Kittaka and Kimura (1989) also ob- Morphologically, the phyllosomas of P. ­echinatus tained success in the culture of phyllosomas of this spe- are very similar to other palinurid species. At stage I the cies feeding pieces of gonads of Mytilus edulis. Dexter mean cephalic length is relatively large compared with (1972) reported that phyllosoma larvae of P. interruptus

Braz. J. Biol., 68(1): 179-186, 2008 185 Abrunhosa, FA., Santiago, AP. and Abrunhosa, JP.

eat large food items: fish larvae, chaetognaths and hy- KITTAKA, J., 1994. Culture of phyllosomas of spiny lobster and dromedusae. In the current study, the phyllosomas were its application to studies of larval recruitment and aquaculture. fed with gonads of mollusk Brachyodonts sp. from instar Crustaceana, vol. 66, no. 3 p. 257-270. III and this kind of food has demonstrated to be suitable KITTAKA, J. and ABRUNHOSA, F., 1997. Characteristics until instar VIII but it was not effective for instar IX be- of palinurids (Decapoda; Crustacea) in larval culture. cause failure moult was observed on many phyllosomas Hydrobiologia, vol. 35, no. 8, p. 305-311. in the tanks. These facts suggest that further research is KITTAKA, J. and IKEGAMI, E., 1988. Culture of palinurid required about the feeding and nutrition of P. echinatus. Panulirus elephas from egg stage to puerulus. Nip. Sui. Gak., Water quality is another decisive factor in the success vol. 54, no. 7, p. 1149 -1154. of the phyllosoma culture. Phyllosomas with good activ- KITTAKA, J., IWAI, M. and YOSHIMURA, M., 1988. Culture ity and free of contaminations by protozoa and bacteria of a hybrid of spiny lobster genus from egg to puerulus depend on rigid control of the water. Any negligence in stage. Nip. Sui. Gak. vol. 54, no. 3, p. 413-417. the water quality can completely affect the larviculture KITTAKA, J. and KIMURA, K., 1989. Culture of the Japanese (Radhakrishnan and Vijayakumaran, 1995). Intensive spiny lobster Panulirus japonicus from egg to juvenile stage. monitoring of the water quality was accomplished dur- Nip. Sui. Gak., vol, 55, no. 6, p. 963-970. ing the culture of P. echinatus. The results of phyllosoma culture may be considered satisfactory compared with INOUE, M., 1978. Studies on the cultured phyllosoma larvae of the , Panulirus japonicus. I. Morphology those reported previously (Kittaka, 1988; Kittaka et al., of the phyllosoma. Nip. Sui. Gak., vol. 44, no. 5, p. 457-475. 1988; Kittaka and Ikegami, 1988). -, 1981. Studies on the cultured phyllosoma larvae of the Japanese spiny lobster, Panulirus japonicus (V. Sievold). 91 p. References (Doctoral Thesis) – University of Tokyo, Tokyo.

ABRUNHOSA, FA., ABRUNHOSA, JP. and COSTA, R. LEWIS, JB., 1951. The phyllosoma larvae of the spiny lobster M., 2004. The early larval stages of phyllosomas of the spiny . Bul. Mar. Sci.Gulf Carib. no. 1, vol. 2, lobsters Panulirus echinatus and P. laevicauda (Decapoda: p. 89‑103. Palinuridae). Nauplius, vol. 12, no. 1, p. 21-29. MATSUDA, H. and YAMAKAWA, T., 2000. The complete BAISRE, JA., 1964. Sobre los estadios larvales de la langosta development and morphological changes of larval Panulirus comun Panulirus argus. Contrib. Cent. Invest. Pesq. Cuba, longipes (Decapoda, Palinuridae) under laboratory conditions. vol. 19, p. 1-37. Fish. Sci., vol. 66, no. 2, p. 278-293. BAISRE, JA. and RUIZ DE QUEVEDO, ME., 1982. Two PHILLIPS, BF. and SASTRY, AN., 1980. The biology and phyllosoma larvae of Panulirus laevicauda (Latreille, 1804) management of lobsters. Academic. New York and London, (Decapoda, Palinuridae) from the Caribbean Sea with a Academic Press. 463 p. discussion about larval groups within the genus. Crustaceana, RADHAKRISHNAN, EV. and VIJAYAKUMARAN, M., 1995. vol. 4, no. 2, p. 147-152. Early larval development of the spiny lobster BOOTH, JD. and PHILLPS, BF., 1994. Early life history of (Linnaeus, 1758) reared in the laboratory. Crustaceana, vol. 68, spiny lobster. Crustaceana, vol. 66, no. 3, p. 271-294. no. 2, p. 151-159. COUTURES, E., 2000. Contributions à l’etude du clicle des SANTIAGO, AP., 2001. Estágios iniciais da lagosta Panulirus Palinuridae et des Scyllaridae (Crustacés Décapodes) du lagon echinatus Smith, 1869 (Decapoda: Palinuridae), cultivados em sud-ouest de Nouvelle-Calédonie 112 p. (Doctoral thesis) – laboratório. 163 p. (Dissertação de Mestrado) – Universidade University of New Caledonia, France. Federal do Ceará, CE. DEXTER, DM., 1972. Molting and growth in laboratory SEKINE, S. SUSUKI, S., SHIMA, Y. and NONAKA, T., reared phyllosomas of the , Panulirus 2000. Larval period and molting in the Japanese spiny lobster interruptus. Calif. Fish. Game, vol. 58, p. 107-115. Panulirus japonicus under laboratory conditions. Fisheries Science. vol. 66, no. 1, p. 19-24. GURNEY, R., 1936. Larvae of Decapoda Crustacea. Part. III. Phyllosoma. Disc. Repor., vol. 12, p. 377-440. YAMAKAWA, T., NISHIMURA, M., MATSUDA H., TSUJIGADO A. and KAMIYA, N., 1989. Complete larval DUGGAN, S. and MCKINNON, AD., 2003. The early larval rearing of the Japanese spiny lobster Panulirus japonicus. Nip. developmental stages of the spiny lobster Sui. Gak., vol. 55, p. 745. (Fabricius, 1798) cultured under laboratory conditions. Crustaceana, vol. 76, p. 313-332.

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